Facilitation by pioneer shrubs for the ecological restoration of riparian forests in the Central Andes of Colombia

Riparian vegetation has great functional importance in agroscapes because it establishes physical connections between natural ecosystems embedded in an agricultural matrix. Throughout the tropics, the historical demand for cropland and pastures in the lands adjacent to streams has led to the replacement of forests by exotic grasses. In order to find new methods for initiating the restoration of riparian forests, we evaluated the use of the pioneer shrubs Tithonia diversifolia and Piper auritum as nurse species for their effects on (1) competition with dominant grasses; (2) natural regeneration and the ecological facilitation of six native tree species of different successional stages; and (3) influence on microclimate. Over a period of 15 months, 4.4% coverage of P. auritum was insufficient to inhibit grasses. In contrast, 81% coverage of T. diversifolia limited the growth of dominant grasses such as Cenchrus purpureus, Paspalum paniculatum, and Cynodon plectostachyus likely by intercepting more than 90% of photosynthetically active radiation (PAR), even though other factors cannot be ruled out. T. diversifolia showed simultaneous effects of facilitation and competition by promoting higher survival of the planted native trees while slowing the growth of pioneer species and inhibiting the regeneration of native woody plants probably as a result of high light interception. This study suggests that planting T. diversifolia as a nurse shrub may facilitate early stages of restoration by inhibiting the growth of pastures, but requires pruning, both to increase light availability and promote the growth of planted trees and the recruitment of woody plants.     

Impacts of litter and understory removal on soil properties in a subtropical Acacia mangium plantation in China

In forest ecosystems, the effects of litter or understory on soil properties are far from being fully understood. We conducted a study in a pure Acacia mangium Willd. plantation in southern China, by removing litter or understory or both components and then comparing these treatments with a control (undisturbed), to evaluate their respective effects on soil physical, chemical and biological properties. In addition, a litter decomposition experiment was conducted to understand the effects of understory on litter decomposition. Our data showed that the presence of understory favored litter decomposition to a large extent. In 1 year, 75.2 and 37.2% of litter were decomposed in the control and understory removal treatment (UR), respectively. Litter had a profound significance in retaining soil water and contributing to soil fertility, including organic matter (OM), available phosphorus (P) and alkali-hydrolyzable nitrogen (N), but understory exerted less influence than litter on soil physical and chemical properties. Both litter and understory played an important role in soil biological activity as indicated by microbial biomass carbon (MBC), while there were no significant impacts on soil exchangeable potassium (K) after either or both were removed. Contrary to our hypothesis, the effects of understory or litter removal were not always negative. A significant soil pH increase with litter removal was a positive factor for acid soil in the studied site. Except for soil moisture, significant effects, caused by removal of litter or/and understory, on measured soil chemical characteristics were only observed in the top 10 cm soil layer, but not in the 10–20 cm layer. Soil available P and exchangeable K contents were significantly higher in the rainy season than in the dry season, however, for the other soil properties, not substantially affected by season.     

Stomatal dynamics and its importance to carbon gain in two rainforest Piper species

The effects of leaf-air vapor pressure deficit (VPD) on the transient and steady-state stomatal responses to photon flux density (PFD) were evaluated in Piper auritum, a pioneer tree, and Piper aequale, a shade tolerant shrub, that are both native to tropical forests at Los Tuxtlas, Veracruz, México. Under constant high-PFD conditions, the stomata of shade-acclimated plants of both species were sensitive to VPD, exhibiting a nearly uniform decrease in g_s as VPD increased. Acclimation of P. auritum to high light increased the stomatal sensitivity to VPD that was sufflcient to cause a reduction in transpiration at high VPD's. At low PFD, where g_s was already reduced, there was little additional absolute change with VPD for any species or growth condition. The stomatal response to 8-min duration lightflecks was strongly modulated by VPD and varied between the species and growth light conditions. In P. aequale shade plants, increased VPD had no effect on the extent of stomatal opening but caused the rate of closure after the lightfleck to be faster. Thus, the overall response to a lightfleck changed from hysteretic (faster opening than closure) to symmetric (similar opening and closing rates). Either high or low VPD caused g_s not to return to the steady-state value present before the lightfleck. At high VPD the value after was considerably less than the value before whereas at low VPD the opposite occurred. Shade-acclimated plants of P. auritum showed only a small g_s response to lightflecks, which was not affected by VPD. Under sunfleck regimes in the understory, the stomatal response of P. aequale at low VPD may function to enhance carbon gain by increasing the induction state. At high VPD, the shift in the response enhances water use efficiency but at the cost of reduced assimilation.     

Photosynthetic light acclimation in two rainforest Piper species with different ecological amplitudes

Summary Piper auritum (H.B. & K.), a pioneer tree restricted to open sites and Piper hispidum (Swartz), a shrub common in sites ranging from recent clearings to shaded understory, both adjust photosynthetic characteristics in response to light availability during growth. The sensitivity of photosynthetic capacity to light availability during growth was indistinguishable for the two species growing in their natural habitat. Photosynthetic capacity was strongly correlated with leaf nitrogen in both species, and the relationship was similar between species. Dark respiration and leaf specific mass were more sensitive to light during growth in P. hispidum, the species with the broad habitat ange, than in P. auritum. In general, similarities between the species were more striking than differences between them. The differences in dark respiration could have important implications for carbon balance. The difference in the responsiveness of leaf specific mass to light indicates that the broad-ranging species maintains access to modes of response little utilized by the open-site specialist. We did not and, in the gas exchange characteristics, any evidence that the open site specialist is better suited than the generalist to high-light sites.This CIW DPB publication number 962     

Removal of invasive shrubs alters light but not leaf litter inputs in a deciduous forest understory

The establishment and spread of non‐native, invasive shrubs in forests poses an important obstacle to natural resource conservation and management. This study assesses the impacts of the physical removal of a complex of woody invasive shrub species on deciduous forest understory resources. We compared leaf litter quantity and quality and understory light transmittance in five pairs of invaded and removal plots in an oak‐dominated suburban mature forest. Removal plots were cleared of all non‐native invasive shrubs. The invasive shrubs were abundant (143,456 stems/ha) and diverse, dominated by species in the genera Ligustrum, Viburnum, Lonicera, and Euonymus. Annual leaf litter biomass and carbon inputs of invaded plots were not different from removal plots due to low leaf litter biomass of invasive shrubs. Invasive shrub litter had higher nitrogen (N) concentrations than native species; however, low biomass of invasive litter led to low N inputs by litter of invasive species compared to native. Light transmittance at the forest floor and at 2 m was lower in invaded plots than in removal plots. We conclude that the removal of the abundant invasive shrubs from a native deciduous forest understory did not alter litter quantity or N inputs, one measure of litter quality, and increased forest understory light availability. More light in the forest understory could facilitate the restoration of forest understory dynamics.     

Holcus lanatus invasion slows decomposition through its interaction with a macroinvertebrate detritivore, Porcellio scaber

Detritus based food webs may mediate the impacts of invasive species on ecosystem processes. Holcus lanatus (L.) is an invasive perennial grass that is rapidly spreading in the coastal prairie of California. We used litterbags to determine if H. lanatus altered the rate of litter decomposition through its interaction with the dominant macroinvertebrate detritivore at our study site, the isopod Porcellio scaber (Latreille). Over the course of a year, H. lanatus litter loss was 15% less than annual grasses, a difference that was directly attributable to P. scaber. Although there was no effect of isopods on litter loss during the winter, when most decomposition occurred, isopods had substantial effects on litter loss of annual grasses during the summer. P. scaber had no effect on litter loss of H. lanatus litter. Our findings suggest that H. lanatus invasion slows decomposition in the coastal prairie because it is unusually refractory to an important detritivore. The decreased decomposition of H. lanatus litter, along with increased production during the growing season, lead to a doubling of the end-of-season litter biomass where H. lanatus had invaded.     

Dynamic stomatal behavior and its role in carbon gain during lightflecks of a gap phase and an understory Piper species acclimated to high and low light

Summary Steady-state and dynamic stomatal and assimilation responses to light transients were characterized in sun- and shade-acclimated plants of Piper auritum, a pioneer tree, and Piper aequale a shade-tolerant shrub from a tropical forest at Los Tuxtlas, Veracruz, México. Despite essentially identical steady-state responses of stomatal conductance to PFD of P. aequale and P. auritum shade plants, the dynamic responses to lightflecks were markedly different and depended on the growth regime. For both species from both growth environments, the increase in stomatal conductance occurring in response to a lightfleck continued long after the lightfleck itself so that the maximum stomatal conductance was not reached until 20–40 min after the lightfleck. Closing then occurred until stomatal conductance returned to near its original value before the lightfleck. Plants that were grown under light regimes similar to those of their natural habitat (high light for P. auritum and shade for P. aequale) had large maximum excursions of stomatal conductance and slower closing than opening responses. Plants grown under the opposite conditions had smaller excursions of stomatal conductance, especially in P. auritum, and more symmetrical opening and closing. The large and hysteretic response of stomatal conductance of P. aequale shade plants to a lightfleck was shown to improve carbon gain during subsequent lightflecks by 30–200%, depending on lightfleck duration. In contrast the very small stomatal response to lightflecks in P. auritum shade plants, resulted in no significant improvement in use of subsequent lightflecks.     

Direct and indirect effects of exotic annual grasses on species composition of a South Australian grassland

Abstract Invasion by Mediterranean annual grasses, such as Avena L. spp. and Bronms L. spp, is one of the major threats to temperate perennial grassland. This study investigated the effects of annual grasses and their litter on the species composition of a grassland near Burra, South Australia. The placement of annual grass litter on soil samples in the glasshouse decreased the establishment or growth of several exotic annual dicots. In the field the addition of annual grass litter slightly decreased the frequency of Danthonia Lam. & DC. tussocks. Furthermore, litter strongly reduced the species richness from 13 species in plots with no litter to nine species in plots with the highest litter level, mainly by decreasing the frequency of common exotic dicots. Native dicot frequency similarly appeared to be decreased by litter addition. In addition to the negative effects of their litter, annual grasses also directly competed with perennial grasses. The magnitude of the competitive effect varied systematically along a slope, suggesting that other factors such as soil properties may control competitive inter actions. The biomass of annual grasses also tended to increase with the addition of their own litter. This combination of positive and negative feedback mechanisms suggests that brief periods favourable for annual grasses, either through management changes or environmental conditions, can lead to persistent changes in the species composition of the system.     

Effects of understory removal, N fertilization, and litter layer removal on soil N cycling in a 13-year-old white spruce plantation infested with Canada bluejoint grass

Canada bluejoint grass [Calamagrostis canadensis (Michx.) Beauv., referred to as bluejoint below] is a competitive understory species widely distributed in the boreal region in North America and builds up a thick litter layer that alters the soil surface microclimate in heavily infested sites. This study examined the effects of understory removal, N fertilization, and litter layer removal on litter decomposition, soil microbial biomass N (MBN), and net N mineralization and nitrification rates in LFH (the sum of organic horizons of litter, partially decomposed litter and humus on the soil surface) and mineral soil (0–10 cm) in a 13-year-old white spruce [Picea glauca (Moench.) Voss] plantation infested with bluejoint in Alberta, Canada. Removal of the understory vegetation and the litter layer together significantly increased soil temperature at 10 cm below the mineral soil surface by 1.7 and 1.3°C in summer 2003 and 2004, respectively, resulting in increased net N mineralization (by 1.09 and 0.14 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004) and net nitrification rates (by 0.10 and 0.20 mg N kg⁻¹ day⁻¹ in LFH and mineral soil, respectively, in 2004). When the understory vegetation was intact, nitrification might have been limited by NH₄ ⁺ availability due to competition for N from bluejoint and other understory species. Litter layer removal increased litter decomposition rate (percentage mass loss per month) from 2.6 to 3.0% after 15 months of incubation. Nitrogen fertilization did not show consistent effects on soil MBN, but increased net N mineralization and nitrification rates as well as available N concentrations in the soil. Clearly, understory removal combined with N fertilization was most effective in increasing rates of litter decomposition, net N mineralization and nitrification, and soil N availability. The management of understory vegetation dominated by bluejoint in the boreal region should consider the strong effects of understory competition and the accumulated litter layer on soil N cycling and the implications for forest management.     

Mixing Effects of Understory Plant Litter on Decomposition and Nutrient Release of Tree Litter in Two Plantations in Northeast China

Understory vegetation plays a crucial role in carbon and nutrient cycling in forest ecosystems; however, it is not clear how understory species affect tree litter decomposition and nutrient dynamics. In this study, we examined the impacts of understory litter on the decomposition and nutrient release of tree litter both in a pine (Pinus sylvestris var. mongolica) and a poplar (Populus × xiaozhuanica) plantation in Northeast China. Leaf litter of tree species, and senesced aboveground materials from two dominant understory species, Artemisia scoparia and Setaria viridis in the pine stand and Elymus villifer and A. sieversiana in the poplar stand, were collected. Mass loss and N and P fluxes of single-species litter and three-species mixtures in each of the two forests were quantified. Data from single-species litterbags were used to generate predicted mass loss and N and P fluxes for the mixed-species litterbags. In the mixture from the pine stand, the observed mass loss and N release did not differ from the predicted value, whereas the observed P release was greater than the predicted value. However, the presence of understory litter decelerated the mass loss and did not affect N and P releases from the pine litter. In the poplar stand, litter mixture presented a positive non-additive effect on litter mass loss and P release, but an addition effect on N release. The presence of understory species accelerated only N release of poplar litter. Moreover, the responses of mass loss and N and P releases of understory litter in the mixtures varied with species in both pine and poplar plantations. Our results suggest that the effects of understory species on tree litter decomposition vary with tree species, and also highlight the importance of understory species in litter decomposition and nutrient cycles in forest ecosystems.     

Do Tree Canopies Enhance Perennial Grass Restoration in California Oak Savannas?

Scattered trees in grass‐dominated ecosystems often act as islands of fertility with important influences on community structure. Despite the potential for these islands to be useful in restoring degraded rangelands, they can also serve as sites for the establishment of fast growing non‐native species. In California oak savannas, native perennial grasses are rare beneath isolated oaks and non‐native annual grasses dominate. To understand the mechanisms generating this pattern, and the potential for restoration of native grasses under oaks, we asked: what are the effects of the tree understory environment, the abundance of a dominant non‐native annual grass (Bromus diandrus), and soils beneath the trees on survival, growth, and reproduction of native perennial grass seedlings? We found oak canopies had a strong positive effect on survival of Stipa pulchra and Poa secunda. Growth and reproduction was enhanced by the canopy for Poa but negatively impacted for Stipa. We also found that Bromus suppressed growth and reproduction in Stipa and Poa, although less so for Stipa. These results suggest the oak understory may enhance survival of restored native perennial grass seedlings. The presence of exotic grasses can also suppress growth of native grasses, although only weakly for Stipa. The current limitation of native grasses to outside the canopy edge is potentially the result of interference from annual grasses under oaks, especially for short‐statured grasses like Poa. Therefore, control of non‐native annual grasses under tree canopies will enhance the establishment of S. pulchra and P. secunda when planted in California oak savannas.     

<Emphasis Type="Italic">Hedychium gardnerianum</Emphasis> Invasion into Hawaiian Montane Rainforest: Interactions Among Litter Quality,Decomposition Rate,and Soil Nitrogen Availability

Few studies have examined the invasion of understory species into closed-canopy forests and, despite inter-specific differences in litter quality and quantity between understory and dominant canopy trees, the influence of understory invasions on soil nitrogen (N) cycling remains unknown. This paper examines litter quality and decomposition of kahili ginger (Hedychium gardnerianum), an invasive understory herb, to determine the influence of this species on N cycling in a Hawaiian montane rainforest. To examine the potential feedback between increased soil N availability and litter decomposition, litter from the invasive ginger, a native tree, and native tree fern was collected from unfertilized and fertilized plots and decomposed in a reciprocal transplant design. Hedychium litter decomposed faster than litter from the two native species. Across species, decomposition rates were negatively correlated with litter lignin content. Despite rapid decomposition rates of Hedychium litter, soil nitrogen availability and rates of net mineralization in the soil were similar in invaded and uninvaded plots. Nitrogen cycling at this site may be more strongly influenced by native species, which contribute the most to overall stand biomass. A negative effect of fertilization on the decomposition of Hedychium litter suggests that a negative feedback between litter quality and soil N availability may exist over longer timescales.     

Interactive effects of elevated CO<Subscript>2</Subscript>, N deposition and climate change on plant litter quality in a California annual grassland

Although global changes can alter ecosystem nutrient dynamics indirectly as a result of their effects on plant litter quality, the interactive effects of global changes on plant litter remain largely unexplored in natural communities. We investigated the effects of elevated CO₂, N deposition, warming and increased precipitation on the composition of organic compounds in plant litter in a fully-factorial experiment conducted in a California annual grassland. While lignin increased within functional groups under elevated CO₂, this effect was attenuated by warming in grasses and by water additions in forbs. CO₂-induced increases in lignin within functional groups also were counteracted by an increase in the relative biomass of forbs, which contained less lignin than grasses. Consequently, there was no net change in the overall lignin content of senesced tissue at the plot level under elevated CO₂. Nitrate additions increased N in both grass and forb litter, although this effect was attenuated by water additions. Relative to changes in N within functional groups, changes in functional group dominance had a minor effect on overall litter N at the plot level. Nitrate additions had the strongest effect on decomposition, increasing lignin losses from Avena litter and interacting with water additions to increase decomposition of litter of other grasses. Increases in lignin that resulted from elevated CO₂ had no effect on decomposition but elevated CO₂ increased N losses from Avena litter. Overall, the interactions among elements of global change were as important as single-factor effects in influencing plant litter chemistry. However, with the exception of variation in N, litter quality had little influence on decomposition over the short term.     

Importance of seed size for the establishment of seedlings of five deciduous broad-leaved tree species

Effects of seed size and phenology on the establishment of five deciduous broad-leaved tree species were examined in deciduous woodland. Treatments included absence and presence of litter in the forest understory, a small gap, and a large gap. Seedling emergence of large-seeded speciesQuercus mongolica var.grosseserrata andAcer mono was not reduced by accumulation of litter in the forest understory, but was promoted in the large gap where litter was less. Seedling emergence of small-seeded species,Alnus hirsuta, Cercidiphyllum japonicum andBetula platyphylla var.japonica, was reduced by the litter in almost all of the sites. Seedlings of large-seeded species avoid shade stress phenologically by unfolding all of their large leaves in a short period before canopy closure in the forest understory. These species had little mortality after seedling emergence. In contrast, small-seeded species have a longer duration of leaf emergence, shorter leaf longevity, and rapid leaf turnover in all the sites. These seedlings attained similar height to those of the large-seeded species at the end of the second year in the large gap, but survival and height growth rate decreased after canopy closure in the forest understory. We suggest that the importance of seed size in determining seedling establishment largely depends on the relationships between seasonal changes of environmental conditions and phenological traits of seedlings, which are related to seed size.Abbreviations Ah Alnus hirsuta - Am Acer mono - Cj Cercidiphyllum japonicum - Bp Betula platyphylla var.japonica - Qm Quercus mongolica var.grosseserrata     

Litter decomposition of a pine plantation is affected by species evenness and soil nitrogen availability

Background and aims

Litter decomposition is a key process controlling flows of energy and nutrients in ecosystems. Altered biodiversity and nutrient availability may affect litter decomposition. However, little is known about the response of litter decomposition to co-occurring changes in species evenness and soil nutrient availability.

Methods

We used a microcosm experiment to evaluate the simultaneous effects of species evenness (two levels), identity of the dominant species (three species) and soil N availability (control and N addition) on litter decomposition in a Mongolian pine (Pinus sylvestris var. mongolica) plantation in Northeast China. Mongolian pine needles and senesced aboveground materials of two dominant understory species (Setaria viridis and Artemisia scoparia) were used for incubation.

Results

Litter evenness, dominant species identity and N addition significantly affected species interaction and litter decomposition. Higher level of species evenness increased the decomposition rate of litter mixtures and decreased the incidence of antagonistic effects. A. scoparia-dominated litter mixtures decomposed faster than P. sylvestris var. mongolica- and S. viridis-dominated litter mixtures. Notably, N addition increased decomposition rate of both single-species litters and litter mixtures, and meanwhile altered the incidence and direction of non-additive effects during decomposition of litter mixtures. The presence of understory species litters stimulated the decomposition rate of pine litters irrespective of N addition, whereas the presence of pine litters suppressed the mass loss of A. scoparia litters. Moreover, N addition weakened the promoting effects of understory species litters on decomposition of pine litters.

Conclusions

Pine litter retarded the decomposition of understory species litters whereas its own decomposition was accelerated in mixtures. Nitrogen addition and understory species evenness altered species interaction through species-specific responses in litter mixtures and thus affected litter decomposition in Mongolian pine forests, which could produce a potential influence on ecosystem C budget and nutrient cycling.     

The population biology of Bromus tectorum in forests: effect of disturbance,grazing, and litter on seedling establishment and reproduction

Summary The effect of tree canopy, understory, herbivores, and litter depth on seedling establishment, survival, and reproduction of the alien grass, Bromus tectorum (cheatgrass), was examined in a series of experiments in four forest habitat types in western North America. Higher recruitment, survival, and reproduction on clearcuts, which would be expected if the overstory alone is limiting the distribution of cheatgrass in forests, were not observed. Removing the understory in an otherwise undisturbed Pinus ponderosa forest did, however, increase the emergence of B. tectorum, but plants in these experimentally-created openings were more vulnerable to grazing by small mammals. In contrast, removing the sparse understory in an Abies forest neither enhanced recruitment nor increased the incidence of grazing of B. tectorum seedlings. Regardless of the forest habitat, most grazed plants died before maturity; even fewer grazed plants produced seeds. Litter depth influenced both recruitment and biomass production: both the rate of germination and the size of resultant seedlings were lower on thick litter (6 cm) compared to results on thin litter (1.5 cm). In the more open Pinus ponderosa and Pseudotsuga menziesii forests, cheatgrass colonization may often occur in openings in the understory alone. Colonization in the more shady A. grandis and Thuja plicata forests is unlikely, however, unless the opening extends through both the understory and the overstory. As a result, cheatgrass is unlikely to increase in any of these forests unless the scale and incidence of disturbance increases substantially.     

The Effects of Exotic Grasses on Litter Decomposition in a Hawaiian Woodland: The Importance of Indirect Effects

Exotic grasses and grass-fueled fires have altered plant species composition in the seasonal submontane woodlands of Hawaii Volcanoes National Park. These changes have altered both structural and functional aspects of the plant community, which could, in turn, have consequences for litter decomposition and nitrogen (N) dynamics. In grass-invaded unburned woodland, grass removal plots within the woodland, and woodland converted to grassland by fire, we compared whole-system fluxes and the contributions of individual species to annual aboveground fine litterfall and litterfall N, and litter mass and net N loss. We assessed the direct contribution of grass biomass to decomposition and N dynamics, and we determined how grasses affected decomposition processes indirectly via effects on native species and alteration of the litter layer microenvironment. Grasses contributed 35% of the total annual aboveground fine litterfall in the invaded woodland. However, total litterfall mass and N were not different between the invaded woodland and the grass removal treatment because of compensation by the native tree Metrosideros polymorpha, which increased litter production by 37% ± 5% when grasses were removed. The 0.3 g N m^–2/y^–1 contained in this production increase was equal to the N contained in grass litter. Litter production and litterfall N was lowest in the grassland due to the loss of native litter inputs. Decomposition of litterfall on an area basis was highest in the grass-invaded woodland. We attributed this effect to increased inherent decomposability of native litter in the presence of grasses because (a) the microenvironment of the three vegetation treatments had little effect on decomposition of common litter types and (b) M. polymorpha litter produced in the invaded woodland decomposed faster than that produced in the grass removal plots due to higher lignin concentrations in the latter than in the former. Area-weighted decomposition was lowest in the grassland due to the absence of native litter inputs. Across all treatments, most litter types immobilized N throughout the incubation, and litter net N loss on an area basis was not different among treatments. Our results support the idea that the effects of a plant species or growth form on decomposition cannot be determined in isolation from the rest of the community or from the direct effects of litter quality and quantity alone. In this dry woodland, exotic grasses significantly altered decomposition processes through indirect effects on the quantity and quality of litter produced by native species.     

Restoring Perennial Warm‐Season Grasses as a Means of Reversing Mesophication of Oak Woodlands in Northern Mississippi

Fire suppression has removed an important ecological force previously responsible for shaping many plant communities throughout the world. Upland areas of north‐central Mississippi that have been protected from fire are now closed‐canopy forests including species known to be uncommon as bearing/witness trees in upland portions of the landscape (historically off‐site species) and sparse ground cover vegetation. Anecdotal evidence suggests that warm‐season grasses were prevalent in the understory of these communities, which could have provided more consistent fuel. We corroborate the historic presence of these grasses by looking at their natural co‐occurrence with oak regeneration (a requisite of self‐replacing stands of oaks found historically). Restoration of these communities has typically focused on burning and off‐site tree thinning. Utilizing a restoration experiment implementing these treatments, we found significantly reduced understory leaf litter in treatment areas. To test which variables associated with restoration treatments were most important for the survival of these grasses, we measured the effect of leaf litter removal and its interaction with environmental conditions on the survival of transplanted shoots. Survival of little bluestem increased with decreasing canopy density and decreasing leaf litter. Leaf‐litter removal did not increase survival, nor did it interact with either pre‐treatment leaf litter depth or canopy density. These results show that little bluestem benefits from conditions expected historically: increased light and possibly fire.     

Dryland management regimes alter forest habitats and understory arthropod communities

Dryland forests, those characterised as having low precipitation and soil nutrients, account for over a quarter of forests globally. Increasing their productivity often relies on irrigation and fertilisation, but the impacts on the wider habitat are largely unknown. Understory invertebrates, in particular, play key roles in forest systems (e.g. nutrient cycling), but their responses to dryland forest management practices are untested. We investigated the impacts of irrigation, fertilisation and a combination of both on soil chemistry, understory vegetation, tree growth and understory arthropod communities in a Eucalyptus plantation to establish linkages between dryland management and ecosystem responses. Fertilisation increased all soil nutrients (N, NO₃N, P and K) with similar effects on the chemical composition of understory grasses. Fertilisation also caused declines in foliar silicon concentrations, an important herbivore defence in grasses. Irrigation increased growth of both understory plants (+90%) and trees (+68%). Irrigation increased the abundance of ground‐dwelling arthropods by over 480% relative to control plots, but depressed higher level taxon arthropod diversity by 15%, declining by a further 7% (?22%) in combined treatment plots. Irrigation also caused a surge in the abundance of Collembola (+1300%) and Isopoda (+323%). Fertilisation drove increases in the abundance of Isopoda (+196%) and Diptera (+63%), whereas fertilisation combined with irrigation increased populations of Thysanoptera (+166%) and Acarina (+328%). Airborne arthropods were less affected, but fertilisation increased the abundance of Apocrita (+95%) and depressed populations of Thysanoptera (?77%). Diptera abundance was positively related to understory vegetation growth, whereas the abundance of other groups (Collembola, Isopoda, Thysanoptera and Acarina) correlated positively with tree growth. We proposed that the large increases in populations of key detritivores, Collembola and Isopoda, were linked to increased leaf litter from enhanced tree growth in irrigated and combined treatment plots. Our findings suggest that dryland management can increase both plant productivity and abundance of arthropods, but cause arthropod diversity at the higher taxon level to decline overall.     

Woody encroachment reduces nutrient limitation and promotes soil carbon sequestration

During the past century, the biomass of woody species has increased in many grassland and savanna ecosystems. As many of these species fix nitrogen symbiotically, they may alter not only soil nitrogen (N) conditions but also those of phosphorus (P). We studied the N‐fixing shrub Dichrostachys cinerea in a mesic savanna in Zambia, quantifying its effects upon pools of soil N, P, and carbon (C), and availabilities of N and P. We also evaluated whether these effects induced feedbacks upon the growth of understory vegetation and encroaching shrubs. Dichrostachys cinerea shrubs increased total N and P pools, as well as resin‐adsorbed N and soil extractable P in the top 10‐cm soil. Shrubs and understory grasses differed in their foliar N and P concentrations along gradients of increasing encroachment, suggesting that they obtained these nutrients in different ways. Thus, grasses probably obtained them mainly from the surface upper soil layers, whereas the shrubs may acquire N through symbiotic fixation and probably obtain some of their P from deeper soil layers. The storage of soil C increased significantly under D. cinerea and was apparently not limited by shortages of either N or P. We conclude that the shrub D. cinerea does not create a negative feedback loop by inducing P‐limiting conditions, probably because it can obtain P from deeper soil layers. Furthermore, C sequestration is not limited by a shortage of N, so that mesic savanna encroached by this species could represent a C sink for several decades.